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Genetic Diversity of Wild, Weedy and Cultivated Forms of Brassica Rapa

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Genetic Diversity of Wild, Weedy and Cultivated Forms of Brassica Rapa Heredity 74 (1995)491—496 Received 30 June 1994 Genetical Society of Great Britain Genetic diversity of wild, weedy and cultivated forms of Brassica rapa J. H. CROUCH, B. G. LEWISt, D. J. LYDIATE AND R. MITHEN* Brass/ca and 0//seeds Research Department, John Innes Centre, Co/ney, Norwich NR4 7UJ and tSchool of Biological Sciences, University of East Anglia, Norwich NR4 7TJ, U.K. RestrictionFragment Length Polymorphisms (RFLPs) were used to study the genetic diversity within and between accessions of 'wild' and cultivated B. rapa. Two of the wild accessions were likely to be escapes from cultivation because of their geographical origins (Argentina and California). The nature of the other three wild accessions (from Turkey, Algeria and Sicily) was not known. Principal components analysis placed the Argentinian, Californian and Turkish accessions within a cluster which contained all the cultivated forms of B. rapa. The other two B. rapa accessions were genetically divergent and, on the basis of their RFLP genotypes, would have been considered to be more distant from the cultivated forms of B. rapa than accessions of B. nigra and B. montana. The implications of these results for germplasm conservation, selection of material for breeding programmes and phylogenetic studies on the origin of Brassica crops are discussed. Keywords:Brassicarapa, diversity, RFLPs, wild germplasm. taxa (Table 1). Five of the wild accessions were inter- Introduction fertile with B. rapa and on the basis of their morph- Recentadvances have been made in Brassica ology, aliphatic glucosinolate content and cytology taxonomy based upon RFLPs. Comparisons between were consistent with being classified as B. rapa L. (R. F. wild and cultivated accessions of B. oleracea L. and B. Mithen and J. H. Crouch, unpublished data). A sixth rapa L. have led to hypotheses concerning the site of 'wild' B. rapa accession from Egypt (UPM 4623) was domestication of Brassica crops and their subsequent found to be B. juncea Coss. & Czern. Of the five wild spread to other geographical regions (Song et al., 1988, accessions, one was collected in Argentina and one in 1992; Warwick & Black, 1991). Wild forms of both B. California. These are very likely to be escapes from oleracea and B. rapa are difficult to distinguish from cultivation as it is highly improbable that wild B. rapa relatively recent escapes from cultivation (Mitchell, occurs in the New World. The other three accessions 1976). This has led to the inaccurate description of were collected from Turkey, Sicily and Algeria. The many accessions in germplasm collections as being accession from Turkey was collected in arable fields 'wild', whereas they are weedy remnants of previous and may also be an escape from cultivation (Oztürk et cultivations. This may lead to incorrect deductions al., 1985). The accession from Sicily was collected concerning the origin of cultivated taxa. It may also from a road verge. No information is available lead to an underestimation of the relative amounts of concerning the origin of the Algerian accession. genetic diversity within and between wild and culti- vated forms of crop species which has implications for Materialsand methods genetic conservation and the choice of genotypes for breeding programmes. Plantswere grown in John Innes No. 1 compost in To examine how incorrect identification of Brassica insect-proof glasshouses in a 18°C day/12°C night accessions may lead to problems with deducing evolu- temperature regime. Supplementary lighting was tionary relationships we obtained from germplasm provided during the winter. Total genomic DNA collections six accessions of B. rapa which were preparations were made from freeze-dried leaf described as 'wild' and compared these with cultivated material of between two and five individuals from each accession. EcoRl-restricted DNA fragments were *Correspondence separated by electrophoresis and were capillary blotted 1995 The Genetical Society of Great Britain. 491 492 J. H. CROUCH ETAL. Table1.Originof Brassica accessions used in the present study Accession Code no. Genome Origin Description and donor number B. rapa subspecies rapifera 99 A Netherlands Stubble turnip oleifera 29 A Germany Oilseed rape cv. Makelsberg oleifera 529 A Canada Oilseedrape cv. Tobin oleifera 532 A Canada Oilseedrape cv. Parkiand italica 189 A Vegetable cv. Brocoletto, HRI 6218 chinensis 333 A Asia Chinese cabbage var. chinesis, IPSR# R-C-26 chinensis 699 A Asia Chinese cabbage cv. Grannaat syl vest ris 73 A Turkey Wild, UPM 6278 sylvestris 74 A Argentina Wild, UPM 5903 sylvestris 75 A Sicily Wild, UPM 6652 sylvestris 76 A Algeria Wild, UPM 6464 sylvestris 79 A California Wild, UPM 1742 Outgroups B. nigra 179 B Greece Ecotype, BGRC 34180 B. nigra 211 B Germany Wild,BGRC 32960 B. montana 196 C France Wild, BOS 12 Germplasm collections prefixes: HRI: Horticulture Research International, UK; IPSR: Institute of Plant Science Reserch, UK; UPM: Universidad Politecnica Madrid, Spain; BGRC: Institut für Pflansenbau, Germany; BOS: Institut National de Ia Recherche Agronomique, France. Table 2. List of probes used in the present study, the Each restriction fragment was considered as a unit number of RFLPs they detected and the RFLP alleles unique character and was used to create a binary matrix. This to either B. rapa subsp. sylvestris no. 75 or no. 76 matrix was used for principal components analyses with the aid of a program provided by M. Ambrose Polymorphic Alleles specific to (John Innes Centre, Norwich) written within the Probe alleles No. 75 No. 76 GENSTAT s software (Rothamsted Experimental Station, Similarity coefficients were calculated pW2E12 8 — 1 England). — according to the Jaccard coefficient (as described in pW9A2 8 1 pW7B6 16 1 1 Gower & Legendre, 1986) for all possible pairwise pW1F6 14 — — comparisons between genotypes (Table 3). The first pW2B7 16 1 1 and second principal components were then graphi- pW2A11 16 — — cally presented (Figs 2 and 3). Similar methods of data analysis have been adopted by Wetton et al. (1987), pOlO 8 — 2 — Song etaL (1990) and Lashermes etal. (1993). p0112 10 1 p0118 7 — 1 pR54 13 — — Results 7 — I pRll5 Onehundred and twenty-three restriction fragments were scored across all accessions. The Brassica accessionstested showed a very high level of poly- morphism (Fig. 1) and no fragments were common for onto a nylon filter. Filters were probed with 11 anony- all accessions. The wild accessions from Sicily and mous gDNA probes derived from libraries of small Algeria contained several unique restriction fragments (0.8—1.6 kb) PstI-restricted fragments from B. napus (Table 2). With the exception of one unique fragment L., B. oleracea and B. rapa. The probes used for which occurred in B. rapa subsp. italica no. 189, none detecting RFLPs are listed in Table 2. Full details of of the cultivated subspecies or the accessions from DNA extraction, blotting and Southern hybridization California, Argentina and Turkey possessed any are provided by Magrath etal. (1994). unique restriction fragments. The Genetical Society of Great Britain, Heredity, 74, 491—496. GENETIC DIVERSITY IN BRASSICA RAPA 493 of Brassica accessions used in the present study Table 3. Simiarity matrix of D values for B. nigra (2,3),B.montana (1,4,31),B.rapa subspp. rapifera (5,10), European oleifera (6,7), Canadian oleifera (8,9),italica(17,18), chinensis (16,19,20)and B.rapa subsp. sylvestris accessions collected from California Description and donor number (11,12), Argentina (13,14,15), Turkey (21,30), Algeria (22,23,24), Sicily (25,26,27,28,29) I Oilseed rape cv. Makelsberg 32522— Oilseedrape cv. Parkiand 421 1 600003—50111 Chinese cabbage var. chinesis, IPSR 7000036— Chinese cabbage cv. Grannaat 9011801103325—0333— 10 0 0 0 0 23 3 32 — 11 01 1 0 33 3 22 2 — 12 0 1 1 0 23 3 23 2 5— 140110233331435—13 01 0 0 23 3 32 2 34— 15 0 0 0 0 33 3 12 1 3 33 3— 16 00 0 1 31 1 1 2 2 22 1 23 — 17 0 0 0 0 12 2 11 2 2 2 1 1 2 1 — 18 00 0 0 22 3 23 2 2 1 1 1 2 1 4 — 19 01 0 0 22 2 12 2 2 1 1 32 2 1 1 — — 20 0 11 0 22 2 23 1 1 1 2 22 2 1 1 5 21 0 11 1 22 2 12 1 22222 2 1 1 2 3 — Germplasm collections prefixes: HRI: Horticulture Research International, UK; IPSR: Institute of Plant Science Reserch, UK; 22 0 0 1 0 10 0 00 0 00 1 1 2 0 1 11 1 2 — UPM: Universidad Politecnica Madrid, Spain; BGRC: Institut für Pflansenbau, Germany; BOS: Institut National de Ia 23 0 1 1 1 1 0 0 00 0 1 0 1 1 1 0 1 11 1 2 8 24 0 0 0 0 10 0 00 0 00 1 0 1 11 0 0 1 2 77— Table 2. List of probes usedEach in restriction the fragmentpresent was consideredstudy, as thea unit 25 0 0 1 1 1 1 0 12 0 1 1 1 1 1 11 11 2 1 232— number of RFLPs they detectedcharacter and and the was usedRFLP to create alleles a binary matrix.unique This 26 0 0 0 1 1 1 0 11 0 1 1 1 1 1 0 0 1 0 1 2 2226— matrix was used for principal components analyses 27 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 0 11 0 1 1 1 1156— 1 1 1 1 0 0 1 1 0 1 1 2 with the aid of a program provided by M. Ambrose 28 0 0 1 0 01 0 11 1 22555— 29 0 0 0 0 12 2 12 0 1 1 22 1 1 0 2 1 2 2 1 105543— 30 0 0 1 0 11 2 11 0 1 2 1 1 1 1 1 1 11 2 11001001— GENSTAT s software (Rothamsted Experimental Station, 31 0 0 0 2 00 0 00 0 0 1 000 1 0 1 1 0 1 000000002— Similarity coefficients were calculated 1234567890123456789012345678901 1 2 3 cally presented (Figs 2 and 3).
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